Temporal resource variability and the habitat-matching rule

Summary The ideal free distribution of competitors in a heterogeneous environment often predicts habitat matching, where the equilibrium number of consumers in a patch is proportional to resource abundance in that patch. We model the interaction between habitat matching and temporal variation in resource abundance. In one patch the rate of resource input follows a Markov chain; a second patch does not vary temporally. We predict patch use by scaling transition rates in the variable patch to the time that consumers require to respond to changes in rates of resource input. If consumers respond very quickly, habitat matching tracks temporal variability. If resource input fluctuates faster than consumers respond, habitat matching averages over the equilibrium of the Markov chain. Tracking and averaging produce the same mean resource consumption for individuals, but long-term mean occupation of the patches differs. When habitat matching tracks temporal variability in resources, consumer density in the variable patch has a lower mean and a higher variance than when habitat matching reflects only average rates of resource input.We tested our model by feeding free-living mallard ducks (Anas platyrynchos) at two artificial patches. The foragers' behavior satisfied the quantitative predictions of the model in each of two experiments.     

Competitor-to-resource ratio, a general formulation of operational sex ratio, as a predictor of competitive aggression in Japanese medaka (Pisces: Oryziidae)

Operational sex ratio (OSR), the number of potentially matingmales divided by the number of fertilizable females, playsa central role in the theory of mating systems by predictingthe intensity of intra-sexual competition and sexual selection.We introduce a general version of OSR, competitor-to-resourceratio (CRR, the number of potential competitors divided bythe number of resource units), as a potential way of predictingthe intensity of competition for any resource. We manipulatedCRR over a broad range (0.5-8) by varying both the number ofcompeting male Japanese medaka fish (Oryzias latipes) and thenumber of resources, either females or food items. We testedwhether the rate of male—male aggression differed dependingon resource type and whether it increased monotonically or followed a dome-shaped relationship with increasing CRR. The patternsof competitive aggression in relation to CRR did not differsignificantly between resource types. In addition, the percapita rate of aggression followed a dome-shaped curve; itwas low when CRR was less than one, initially increased as CRR increased, was highest at a CRR of about two, and then decreasedwhen CRR was greater than two. However, competitor number,independent of CRR, had a significant and negative effect onrate of aggression. We suggest that CRR is a valuable predictorof the rate of competitive aggression and may be a useful conceptfor synthesizing ideas about resource competition and monopolization that are currently dispersed in the separate bodies of literatureon mating systems, social foraging and territoriality.     

Simultaneous changes in cell quotas of microcystin, chlorophyll a, protein and carbohydrate during different growth phases of a batch culture experiment with Microcystis aeruginosa

The cell quotas of microcystin (Q_mcyst), protein (Q_prot), chlorophylla (Q_chloro) and carbohydrate (Q_carbo), as well as the net productionrates of these parameters, were determined during the exponentialand stationary phases in nine batch cultures of Microcystisaeruginosa (CYA 228) at light regimes from 33 to 53 µmolphotons m^–2 s^–1. The following results were obtained.(i) A parallel pattern was found in the changes of Q_mcyst, Q_prot,Q_chloro and Q_carbo during the entire growth cycle and significantcorrelations were recorded between Q_mcyst and Q_prot, Q_chloroand Q_carbo. (ii) The net microcystin production rate (µ_mcyst)was positively correlated with the specific cell division rate(µ_c), the chlorophyll production rate (µ_chloro)and the protein production rate. (iii) A significant inverselinear relationship was found between µ_c and Q_mcyst, i.e.cultures with a positive µ_c had a Q_mcyst between 110 and400 fg microcystin cell^–1, while declining cultures hadQ_mcyst values >400 fg microcystin cell^–1. Maximum variationin Q_mcyst within cultures was 3.5-fold. Collectively, the resultsshow that cells produced microcystin at rates approximatingthose needed to replace losses to daughter cells during divisionand that microcystin was produced in a similar way to proteinand chlorophyll, indicating a constitutive microcystin production.     

Is the between-population variance negligible in the total variance of heterozygosity? Case of a finite number of loci subject to the infinite-allele model in finite monoecious populations

The decomposition of the variance of the average heterozygosity into variances between and within populations is studied in the general case of a finite number of loci. These loci are assumed randomly distributed over chromosome pairs having a non-interference recombination scheme, and independently subject to mutation according to the infinite-allele model. The equilibrium behavior of that decomposition is discussed in the monoecious mating case with regard to each parameter of the model: mutation rate per gene per generation (u), population size (N), number of loci (n), map length of chomosome pairs (L). It is shown that the proportion Q of the between-population variability in the total variance of the average heterozygosity is decreasing as either the mean heterozygosity (θ = 4Nu/(1 + 4Nu)) or the mean number of mutations per gamete per generation (v = nu) is increasing. Moreover, even if Q is always smaller than for this model, it is not negligible unless θ is close to one or v is much larger than one for L long enough.     

Female-specific resource limitation does not make the opportunity for selection more female biased

Competition for limiting resources and stress can magnify variance in fitness and therefore selection. But even in a common environment, the strength of selection can differ across the sexes, as their fitness is often limited by different factors. Indeed, most taxa show stronger selection in males, a bias often ascribed to intense competition for access to mating partners. This sex bias could reverberate on many aspects of evolution, from speed of adaptation to genome evolution. It is unclear, however, whether stronger opportunity for selection in males is a pattern robust to sex-specific stress or resource limitation. We test this in the model species Callosobruchus maculatus by comparing female and male opportunity for selection (i) with and without limitation of quality oviposition sites, and (ii) under delayed age at oviposition. Decreasing the abundance of the resource key to females or increasing their reproductive age was challenging, as shown by a reduction in mean fitness, but opportunity for selection remained stronger in males across all treatments, and even more so when oviposition sites were limiting. This suggests that males remain the more variable sex independent of context, and that the opportunity for selection through males is indirectly affected by female-specific resource limitation.     

An Experimental Approach to the Effects of Predictability on Community Structure

This paper considers the effects of different levels of resourcepredictability on niche widths, competition and diversity ina community of three hummingbird species. Three pairs of hypothesesare tested: whether decreasing predictability (a) increasesor decreases niche width, (b) increases or decreases competitionand (c) increases or decreases diversity. The results show thatcomplementary increases and decreases in niche width occur withdecreasing predictability, while competition and diversity decrease,at least with extreme unpredictability. A model, which assumesthe species to have similar resource preferences, and its predictionsare examined. The dominant species, Lampornis clemenciae, excludes a subordinatespecies, Archilochus alexandri, from preferred resources. Thisdefense becomes unprofitable with decreasing predictabilityand Archilochus invades the resources once vigorously defendedby Lampornis. When the spectrum of resources is altered in thedirection of decreasing suitability for both species, Lampornisbecomes even more specialized and sensitive to the effects ofunpredictability. A third species, Eugenes fulgens, steals resourcesundefended by Lampornis. These characteristics seem to be commonto a number of communities in very different taxonomic groups,and characteristic of communities in which species share a commonpreferred resource. The question of how a community organizedwith distinct resource preferences responds to decreasing predictabilityremains open.     

A Test for the Resource Remobilization Hypothesis: Tree Sprouting using Carbohydrates from Above-ground Parts

To test the resource remobilization hypothesis, i.e. the hypothesisthat some trees sprout from root-collars or from the lower partof trunks using resources obtained from above-ground parts ratherthan from resources reserved in their roots, we conducted cuttingexperiments forEuptelea polyandra, a frequently sprouting treespecies with little carbohydrate reserves in its roots,Quercusserrata,a frequently sprouting tree species with large reservesin the roots, andMallotus japonicus, a rarely sprouting treespecies. Trees of each species were cut down in winter leavingtwo kinds of stumps, those approx. 1.5 m in height and thosecut off near the ground. The number and total dry weight ofnewly sprouted shoots per stump were compared between the twotreatments and among the three species at the end of the followinggrowing season. InE. polyandra,both the number and total dryweight of sprouts per stump were very small for both treatmentsand were similar to, or less than, those ofM. japonicus. Onthe other hand,Q. serratasprouted abundantly in both treatments.These results indicate thatE. polyandracannot sprout sufficientlywithout a considerably large volume of above-ground parts orthat additional structures such as foliage and branches maybe necessary for sprouting. We conclude that the resource remobilizationhypothesis is supported for this species.Copyright 1998 Annalsof Botany Company Euptelea polyandraSieb. et Zacc,Quercus serrataThunb,Mallotus japonicus(Thunb.) Muell. Arg., tree sprouting, cutting experiment, resprouter, resource movement, carbohydrate allocation, ground-surface disturbance, root stock, resource remobilization hypothesis.     

Mechanisms and fitness effects of interspecific information use between migrant and resident birds

Interactions with potential competitors are an important componentof habitat quality. Due to the costs of coexistence with competitors,a breeding habitat selection strategy that avoids competitorsis expected to be favored. However, many migratory birds appearto gain benefits from an attraction to the presence of residentbirds, even though residents are assumed to be competitivelydominant. Thus far the mechanisms of this habitat selectionprocess, heterospecific attraction, are unknown, and the consequencesfor resident birds of migrant attraction remain untested. Throughheterospecific attraction, migrants may gain benefits if thedensity or territory location of residents positively reflectshabitat quality, and/or they gain benefits through increasedfrequency of social interactions with residents in foragingor predator detection. In this experiment, we examined the reciprocaleffects of spatial proximity on fitness-related traits in migrantpied flycatcher (Ficedula hypoleuca) and resident great tit(Parus major) by experimentally forcing them to breed eitheralone or in close proximity to each other. Surprisingly, greattits bore all the costs of coexistence while flycatchers wereunaffected, even gaining slight benefits. In concert with anearlier study, these results suggest that flycatchers use titsas information about good-quality nest-site locations whilebenefits from social interactions with tits are possible butless important. We suggest that utilizing interspecific socialinformation may be a common phenomenon between species sharingsimilar resource needs. Our results imply that the effects ofinterspecific information use can be asymmetric and may thereforehave implications for the patterns and consequences of speciescoexistence.     

Foraging innovation is inversely related to competitive ability in male but not in female guppies

Foraging success is likely to affect hunger level and motivationto locate and exploit novel food sources in animals. We exploredthe relationship between scramble competition for limited foodand foraging innovation in the guppy (Poecilia reticulata),predicting that poor competitors would be more likely to innovatewhen presented with novel foraging tasks. Among males, we foundthat latency to complete novel foraging tasks was correlated bothwith weight gain and number of food items consumed, suggestingthat poor competitors are more likely to innovate. However,among females there was no relationship between innovative tendencyand either weight gain or foraging success. We suggest thatthis sex difference may reflect parental investment asymmetriesin males and females, and we predict similar sex differencesin other species.     

Mechanism of Electron Flow through the QB Site in Photosystem II. 1. Kinetics of the Reduction of Electron Acceptors at the QB and Plastoquinone Sites in Photosystem II Particles from the Cyanobacterium Synechococcus vulcanus

Various derivatives of benzoquinone (BQ) were found to be reducedat two sites [the Q_B and plastoquinone (PQ) sites] in photosystemII particles from Synechococcus vulcanus, and the relationshipbetween the structures of BQs and the kinetics of such reductionat each site were studied. Affinities of BQs for both the Q_B and the PQ site and the maximumturnover rates at the two sites were estimated by computer simulationof the dependence on the concentration of the BQ of the rateof oxygen evolution. Affinities of BQs for the Q_B sites determinedby this method agreed well with those determined from competitionbetween BQs and DCMU for the Q_B site. All methyl-substituted BQs had low affinities for the Q_B site,and tetramethyl-p-benzoquinone had almost no affinity. An increasein the number of chlorine atoms in the quinone ring increasedthe affinity, and the position of such substitutions had a greateffect on the affinity when two positions on the ring were occupiedby two chlorine atoms or methyl groups. The affinities of BQs for the PQ site were almost the same forall BQs tested in this experiment but the maximum turnover ratesat this site varied greatly from one derivative to another. The results are consistent with the hypothesis that the bindingof the PQ molecule to the Q_B site is attributable not to itsquinone ring but to its isoprenoid chain and, moreover, thatthe electron transfer through the Q_B site occurs not by replacementof the PQ molecule but by donation of electrons to another PQmolecule. (Received September 20, 1994; Accepted February 27, 1995)     

Mating system, sexual dimorphism, and the opportunity for sexual selection in a territorial ungulate

In mammals, species with high sexual size dimorphism tend tohave highly polygynous mating systems associated with high variancein male lifetime reproductive success (LRS), leading to a highopportunity for sexual selection. However, little informationis available for species with weak sexual size dimorphism. Ina long-term study population, we used parentage analysis basedon 21 microsatellite markers to describe, for the first time,variance in male lifetime breeding success (LBS) of roe deer,a territorial ungulate where males weigh less than 10% morethan females. LBS ranged from 0 to 14 (mean = 4.54, variance= 15.5), and its distribution was highly skewed, with only afew males obtaining high LBS and many males failing to breedor siring only one fawn. As predicted for polygynous specieswith low sexual size dimorphism, the standardized variance inmale LBS was low (I_m = 0.75) and was only slightly higher thanthe standardized variance in female LRS (I_f = 0.53), suggestinga low opportunity for sexual selection. The I_m value reportedhere for roe deer is much lower than values reported for highlydimorphic ungulates such as red deer (I_m > 3). We suggestthat, along a continuum of opportunity for sexual selection,roe deer occupy a position closer to monogamous and monomorphicterritorial ungulates than to highly polygynous, sexually dimorphicungulates with dominance rank–based mating systems suchas harems or roving mating systems.     

Summer coastal zooplankton biomass and copepod community structure near the Italian Terra Nova Base (Terra Nova Bay, Ross Sea, Antarctica)

The structure of the zooplankton biotic community and of copepodpopulation in the coastal area of Terra Nova Bay (Ross Sea,Antarctica) was investigated during the 10th Italian AntarcticExpedition (1994/1995). Zooplankton biotic community consistedmainly of pteropods (Limacina helicina and Clione antarctica),Cyclopoid (Oithona similis), Poecilostomatoid (Oncaea curvata)and Calanoid (Ctenocalanus vanus, Paraeuchaeta antarctica, Metridiagerlachei and Stephos longipes) copepods, ostracods, larvalpolychaetes and larval euphausiids. Zooplankton abundance rangedfrom 48.1 ind m^–3 to 5968.9 ind m^–3, and copepodabundance ranged from 45.2 ind m^–3 to 3965.3 ind m^–3.The highest peak of zooplankton abundance was observed between25 m and the surface and was mainly due to the contributionof O. similis, O. curvata and C. vanus. Zooplankton biomassranged from 5.28 mg m^–3 to 13.04 mg m^–3 dry weight;the maximum value was observed between 25 m and the surface.Total lipid content varied from 216.44 to 460.73 mg g^–1dry weight.     

The role of resource dispersion in promoting the co‐occurrence of dominant and subordinate ant species

Dominant competitors govern resource use in many communities, leading to predictions of local exclusion and lower species diversity where dominant species are abundant. However, subordinate and dominant species frequently co‐occur. One mechanism that could facilitate resource sharing and co‐occurrence of dominant and subordinate competitors is fine‐scale resource dispersion. Here, we distributed 6 g of a food resource into 1, 2, 8, 32 or 64 units in small 0.40 m² areas centred on nests of the dominant ant Monomorium sydneyense. We tested three hypotheses. First, we hypothesized that the species richness and abundance of foraging ants would increase with increasing resource dispersion. Accordingly, species richness doubled and total ant abundance was two orders of magnitude higher in high resource dispersion treatments. Secondly, we hypothesized that increasing resource dispersion would reduce competitive interactions such as resource turnover events and lower the probability of food resources being occupied. Substantial support for this hypothesis was observed. Finally, we tested the hypothesis that the foraging time of each species would be proportional to the relative abundance of each species solely in high resource dispersion treatments. Expected and observed foraging times were statistically similar for only the dominant ant M. sydneyense. The subdominant Pheidole rugosula increased its foraging time much more than was expected, while two subordinate ants showed no relationship between observed and expected times. Thus, while increasing resource dispersion significantly increased overall species richness, this increase in co‐occurrence did not correlate with a significant increase in foraging time for the two subordinate species. Rather, changes in resource dispersion appeared to benefit only the subdominant species. Inter‐site variation appeared more important for other subordinate species indetermining co‐occurrence and foraging time. Multiple mechanisms facilitate co‐occurrence and resource sharing in this community, and probably in most other communities.     

Comparing measures of breeding inequality and opportunity for selection with sexual selection on a quantitative character in bighorn rams

The reliability and consistency of the many measures proposed to quantify sexual selection have been questioned for decades. Realized selection on quantitative characters measured by the selection differential i was approximated by metrics based on variance in breeding success, using either the opportunity for sexual selection I_s or indices of inequality. There is no consensus about which metric best approximates realized selection on sexual characters. Recently, the opportunity for selection on character mean OSM was proposed to quantify the maximum potential selection on characters. Using 21 years of data on bighorn sheep (Ovis canadensis), we investigated the correlations between seven indices of inequality, I_s, OSM and i on horn length of males. Bighorn sheep are ideal for this comparison because they are highly polygynous and sexually dimorphic, ram horn length is under strong sexual selection, and we have detailed knowledge of individual breeding success. Different metrics provided conflicting information, potentially leading to spurious conclusions about selection patterns. I_δ, an index of breeding inequality, and, to a lesser extent, I_s showed the highest correlation with i on horn length, suggesting that these indices document breeding inequality in a selection context. OSM on horn length was strongly correlated with i, I_s and indices of inequality. By integrating information on both realized sexual selection and breeding inequality, OSM appeared to be the best proxy of sexual selection and may be best suited to explore its ecological bases.     

Estimation of sample zooplankton abundance from Folsom splitter sub-samples

Procedures are described for estimating the abundance of eachzooplanklon species in a sample after counting at least twosub-samples from a Folsom splitter. A multinomiai model is assumedfor the splitting process. The method may be summarized as follows:two sub-samples, balanced with respect to the left — rightsplits, are counted. The counts for each species are testedfor homogeneity using a x² test. If the counts are homogeneous,an estimator is given which permits the estimation of the numberof animals in the sample, and the variance of this estimate.If the subsample counts are heterogeneous, it is assumed thatclumping has occurred. A procedure is described in which additionalsub-samples are counted to locate the clump, and estimatorsof abundance (and variance estimators) are derived. If the splitterbias is <5%, bias of the abundance estimator is negligiblecompared to the binomial sampling error. Comparisons of speciesabundance between plankton samples is made more rigorous usingthe estimates of mean abundance, and the variance of this mean,provided by the methods described here. ^*Present address: Dunoon Road, Dorroughby, NSW 2480, Australia.     

Resource quality effects on Daphnia longispina offspring fitness

The present study addressed the question: is the fitness ofDaphnia longispina neonates related to the quality (species)of the resource their mothers consumed? A 2 2 factorial designwith two maternal and neonate resource types was employed, withRhodomonas and Microcystis used as high- and low-quality resources,respectively. Neonate fitness was assessed using demographicparameters obtained from a life-table study. The resource typeneonates consumed always had a stronger effect on the neonatesthan did the maternal type. Neonate consumption of Rhodomonas.relative to Microcystis. increased fitness, as estimated bythe instantaneous population growth rate, by 25–28%. whilematernal consumption of Rhodomonas increased fitness by 6-8%.Neonate Rhodomonas consumption also reduced the age at reachingthe primiparous instar; increased brood size at the primiparousinstar and body length immediately after the primiparous instar;increases average dutch size, number of clutches produced, totalegg production. and mean body length up to and on age 30 days.Regardless of resource type, maternal Rhodomonas consumptionresulted in neonates which reached maturity at an earlier ageand had a larger individual size immediately after reachingmaturity. The mean clutch size, total egg production, and lengthup to and at age 30 days were all significantly affected bymaternal diet when neonates consumed Microcystis; however, thesevariables were not influenced by maternal type when neonatesconsumed Rhodomonas. This is the first study to document maternallymediated effects of resource quality on the fitness of crustaceanzooplankton neonates.     

Attributes of Individual Flowers of Brassica napus L. are Affected by Defoliation but not by Intraspecific Competition

To investigate the effect of intraspecific competition on floweringin Brassica napus‘Westar’, a series of 30 pots wasestablished that spanned a range of one–96 plants perpot. In each pot, the following attributes of individual flowerswere quantified: petal length; petal width; stamen length; pistillength; pollen production; diameter of pollen grains; and nectarproduction. Certain plants contained a transgenic marker thatenabled the vigour of their pollen to be quantified by screeningthe progeny from post-pollination trials against conventionalmale competitors. Plant size was strongly affected by plantdensity; plants in the lowest density pots comprised ten-timesmore dry biomass than individuals in the highest density pots.However, none of the attributes of individual flowers variedwith density. In contrast, the number of flowers per plant declinedrapidly with density. In the face of resource scarcity, theplants apparently conserved flower size rather than flower number.There was no difference in the response to density between transgenicand conventional plants, but transgenic plants obtained morefertilizations than expected in post-pollination competitionagainst conventional competitors. A separate experiment demonstratedthat B. napus is, nevertheless, capable of plasticity in certainflower attributes (petal size, stamen length) in response todefoliation. Explanations for the stability of flower size relativeto flower number are discussed. Copyright 2001 Annals of BotanyCompany Brassica napus, defoliation, density effects, flowers, flower size, intraspecific competition, nectar production, oilseed rape, pollination, transgenic plants     

Water-column chlorophyll in an oligotrophic environment: correction for the sampling depths and variations of the vertical structure of density, and observation of a growth period

The chlorophyll content of a water column (WCC), which is commonlyused as an index of the phytoplankton abundance, is affectedby the choice of the sampling depths and by the variations ofthe vertical structure of density. For instance, the thicknessof the water layer, between two sigma-t values, which containsthe deep chlorophyll maximum, can vary with internal waves.The resulting noise often dominates the mesoscale variationsof the observed water-column chlorophyll (OWCC). Sigma-t dependentstatistics (mean, standard deviation) of the chlorophyll concentrationare computed using the observations at 29 casts from a 22-day-longfixed station in an oligotrophic environment at 15•S, 173°E.For each cast, these statistics, the sampling depths, and thewater density at these sampling depths, allow the estimationof a station-dependent ‘expected water-column chlorophyll’(EWCC). The ratio of EWCC to the overall likelihood of WCC duringthe fixed station (i.e. the mean of all OWCC) is a measure ofthe effect of sampling and variable density structure at eachcast. When this effect is removed, the noise in WCC estimatesdecreases significantly. The time variations of WCC during thefixed station then show a trend with relatively high valuesduring the first days, followed by a 12-day-long period withlow values. A regular increase occurred from 1 October, whichwas accompanied by high carbon fixation rates and was mainlydue to an increase of the chlorophyll concentration betweenthe surface and the deep chlorophyll maximum. New productionduring this active phase was estimated to be 535 mgC m^–2day^–1, corresponding to 62% of the total production. Breakingof internal waves which were recorded at the beginning of thegrowth phase and vertical mixing of nutrients can explain theobservation.     

Mesozooplankton community structure and grazing impact in the region of the Subtropical Convergence south of Africa

Mesozooplankton distribution and community structure in theregion of the Subtropical Convergence (STC) south of Africawere investigated during the SAAMES III cruise in austral winter(June–July) 1993. Both the STC and an associated warm-coreeddy (WCE) exhibited enhancements in zooplankton abundance,compared to the Subantarctic waters. Particularly, elevatedzooplankton densities were found in the centre of the STC andin the region north of it as well as at the edge of the WCE.Copepods (mainly Pleuromamma abdominalis and Metridia lucens),euphausiids (Euphausia spinifera, E.similis and E.recurva),pteropods (Limacina spp.) and chaetognaths (Eukrohnia hamataand Sagitta spp.) dominated numerically and accounted for >60     

Development of Sinks as an Autocatalytic Feed-back Process: a Test using the Asymmetric Growth of Leaves in Mestha (Hibiscus cannabinus L.)

Differential development of sinks that depend on a common resourcepool has been viewed as a consequence of an autocatalytic feedbackprocess of flow of resource units into them. The feed-back processimplies that the stronger a sink is relative to its competitors,the greater is its probability of getting further resourcesas a non-linear function of its resource drawing ability andsink size. We show that this model contrasts with that of thesink-strength dependent model in its prediction of the subsequentdevelopment of the initial asymmetry of growing leaves whentheir resource drawing ability is enhanced. By artificiallyenhancing the resource drawing ability of the leaves of Mestha(Hibiscus cannabinus L.) by external application of growth regulators,we test these predictions and show that the results are in conformitywith the autocatalytic model proposed by Ganeshaiah and UmaShaanker.Copyright 1995, 1999 Academic Press Autocatalytic growth, resource flow, leaf asymmetry, sink-strength, self-organization, Hibiscus cannabinus (L.)